Torrey Botanical Society

Studies on the Nutritive Value of Spartina Grass Growing in the Marsh Areas of Coastal Georgia

Author(s): Paul R. Burkholder

Source: Bulletin of the Torrey Botanical Club, Vol. 83, No. 5 (Sep. - Oct., 1956), pp. 327-334
Published by: Torrey Botanical Society
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BULLE T IN O F THE T ORR EY B O T A NI C A L C L UB
VOL. 83, No. 5, pp. 327-334 SEPTEMIBER, 1956

STUDIES ON THE NUTRITIVE VALUE OF SPARTINA GRASS

GROWING IN THE MARSH AREAS OF COASTAL GEORGIA1
PAUL R. BURKHOLDER2

Extensive low areas along the coast of Georgia are coveredwith marsh
grasses. Among these,Spartina alterniflora,Loisel. contributeson a large
scale to primaryproductivityin the marshesand shallow estuaries of the
region. Especially luxuriant growth of this dark green grass, oftentimes
attaining a height of six feet, occurs in the low niarshes, along river banks
and on the shores of islands. The grass of high marshland is yellow green
in color, tougher, aind generally less thani two feet in height. The Spartiva
main crop grows during a lolngseason frolmIMarch to October. In protected
places with southern exposure, however, the grass may show growth of
telider green shoots even during the wiTinter months. The crop apparently
disilntegratesslowly during the year or two following its mllaturationin the
fall. Microbial conversion of grass and its consumption by small herbivores
leads illto the conmplexcycle of nutritioniwhich suLpportscrustaceans, fish,
and fowl, all of which aboulnd in the regioln.
Duriiig spring and early suniiiier, cattle are known to graze on the
salt marshes where Spartina is aceessible. The grass is harvested in silall
quantities, for cattle feeding, by residents of the sea islands. The writer
was told by Mr. Frank Durand, who resides o1n Sapelo Islalnd, that in
fornmeryears marsh grass was harvested as the sole feed for ilules that
were used to haul fuel for the wood-burning steain locomeotiveson the old
Bruniswick and Florida Railroad (Black 1952).
General observations of the extensive productivity aind potential uses
of Spartina in the coastal regions of Gleorgia prompted us to collect sam-
ples of the inarsh grass for analysis of its niutritivecontent, with particular
reference to its B vitamins and amiino aeids, known to be essential for grow-
ing animals.
Materials and methods. Four differenitsamples of imiaturemarsh grass
were gathered from different locations, along river banks and on high
marsh in the vicinity of Sapelo Island in the period August 14 to 27, 1955,
and dried rapidly with heat supplied by 200-watt incandescent lamps.
Young grass was harvested from a southern exposure on the bank of
1 These studies were aided by a contract between the Officeof Naval Research, De-
partmentof the Navy, and the University of Georgia, NR163 308. Some of the work was
done at the Marine Biology Laboratory, Sapelo Island, Georgia, and this paper is con-
tribution No. 3 from that laboratory.
2 The author's present address is Brooklyn Botanic Garden, Brooklyn 25, N.Y.

327
[THE BULLETIN' for July-August, 1956 (83: 253-325) was issued August 20, 1956]

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328 BULLETIN OF THE TORREY BOTANICAL CLUB [VOL. 83

Lazaretta Creek, near Savannah, Georgia, on Novemuber29, 1955, and

dried withheat froma steamradiator.Dead grass stemsfromthe preceding
year's growthwere gathered on August 27, 1955 from a pile left by a
spring tide. These old stems were devoid of leaves, whieh long ago had
disintegratedand washed away. The stemswere dried over a steamradiator.
A summaryof the data eolneernilngsoureematerialsis presentedin Table 1.
TABLE 1. Dcscription of grass samiples collected for chtemicaland inicrobiological
analyses.

- ~ ~ ~ ~ ~ ~ ~ ~

I Mature Terminial Leaf- Low Marsh, August 14, 1955 Tungsteii

stalks, 2 ft. long J-aekCreek lamps 1.5.1
2 Mature Terminal Leaf - High Marsh, August 20, 1955 Tungsten
stalks, I ft. long Bai-iiCreek lamps 14.2
3 Mature Terminal Leaf- Lo++,Marsh, August 22, 1955 Tuiigsten
stalks, 2 ft. long MAud Creek lamps 15.,7
4 Mature Whole plants, Low Marsh, Nugust 27, 1955 Tungstei-i
leaves and stems, Jack Creek lamps 15.0
4-5 ft. long
6 Young Termiiial shoots. Low Marsh, November29, 1955 Steam
About 11/2 ft. long Lazaretta,Creek radiator 13.1
7 Weathered stems Low Marsh, August 27, 1955 Steam
Duplin River radiator 8.7

The dried grass samples were prepared for analyses by grilndingin a

Wiley mill. All samples were kept in cardboard cartons until they could
be analyzed. The moisturecontentwas determined,prior to making the
analyses, so that appropriate correctionscould be made on ali oven dry
basis. Moisture content of the samples, determinedat the time when
proximate analyses were made, is shown in Table 1. The per celit of
moistureat the time when vitaminsand amino acids were determinedwas
found to be as follows: sample no. one, 6.13; sample no. three,6.38; and
sample no. six, 8.77.
Determinationsof moisture,fat, protein,crude -fibre,ash, nitroo-eii-free
extract,calcium,phosphorus,and iron were made accordingto the methods
recommendedin the 8th edition of the OfficialMethods of Analysis of the
Associationof OfficialAgricultural Chemists (1955). These analyses were
made, on six differentsamples, by LJawand Company,Atlanta, Georgia.
The data have been calculated on a dry basis for the reports in Table 2
and Table 3.

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1956] BURKHOLDER: NUTRITIVE VALUE OF SPARTINA GRASS 329

Three samplesof grass were analyzed fortheircontentof ten B vitamins

and ten essential amino acids by Food Research Laboratories,Long Island
City, New York. The techniquesrecommendedin Methods of Analysis of
the Associatioll of OfficialAgricultural Chemists (1955) were employed
for the determillationof thiamine,riboflavin,niacin, total folic acid, and
vitaminB12 activity.Total pantothenicacid was assayed according to the
procedure of Toepfer, Zook and Richardsoii (1954). Biotin and choline
were determiniedaccording to the method outlined by Barton-Wright
(1952). Pyridoxine was assayed by the technique of Parrish, Loy and
Kline (1955). For the assay of inositol,the sample was hydrolyzedby the
method of Jurist and Foy (1944), and the determinationwas made ac-
cording to the general procedure of Atkin, Schultz, Williams, and Frey
TABLE 2. Proximate analyses of different samples of marsh grass and Coastal
Bermuda Grass.3 Values are given as per cent of dry weight, and energy as thermsper
lb.

Samples
1 2 3 4 6 7

Analyses Oe

Ql >t CZ Qt X t CZ

Fat 2.80 2.39 2.39 2.15 2.98 0.78 2.0

Protein 8.51 5.68 9.78 7.86 13.24 4.04 13.1
Crude Fibre 29.38 27.87 31.01 31.20 29.75 35.63 33.0
Ash 9.84 11.72 11.48 10.63 12.83 13.85 4.6
Nitrogen-free
Extract 49.47 52.34 45.34 48.16 40.20 45.70 47.3
Therms 1.70 1.63 1.64 1.69 1.68 1.68

3 Data for Coastal Bermuda Grass (Burton 1948) are shown in the last column for
comparison.

(1943). With the exception of the thiochromemethod for determination

of thiamine,all of the methodsfor assay of the B vitamins were micro-
biological. The resultingdata are reported in Table 4 as milligramsper
100 grams of dry weight.
The ten amino acids were determinedon properlyhydrolyzedsamples
by means of microbiologicalproceduressummarizedin the book by Barton-
Wright (1952). The data, presentedin Table 5, show the per cent of each
amino acid on a dry weightbasis, and also the per cent of each compound
calculated on the basis of nitrogentaken to be 16. This latter expression
gives an approximate per cent of individual amino acids in the mixed
proteinsof the dried grass.
Discussion of data. Proximate analyses, given in Table 2, show minor
variations among the differentsamples of grass. Most interestingis the

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33013 BIU-LLETIN OF THE TORREY BOTANICAL CLUB [FaaL. 83

high per cent of protein, 13.24, in rapidly growing younigleafy shoots

of sample no. 6. The weatheredstems,devoid of leaves after a year, show
loss of fat and protein resultingfrom processes of disintegration.Crude
fibreseems to persist in the weathered steimis, that float and somnetimes
remain on the beaches for a long while. Conmparisonof the data for
Spartina and proximateanalyses made on Coastal Bermuda Grass (Burton
1948), shown in the last column of Table 2, indicate a similarityin com-
position. All of the Spartina samples, however,show a higher contentof
ash, probablybecause of theirsea water habitat. Determinationsof therms
per pound by colorimetryyield data similar to the values obtained by
calculation,on the basis of 4 calories per gmi.of protein or carbohydrate,
and 9 ealories per gm. of fat.
TABLE 3. Somie miiineral
valutesof Spartinia expres,nsed
(Ia. per cenltof each elemitentin
the dried grass.

Samples
1 2 3 4 6
Mlillerals Matuie MatuIre Mattlile re.i
ves You'lng Aveathei(ld

Leaves Leaves Leaves L eaves

aiid Stems Ste'lis

Ca 1.03 0.71 1.47 0.39 0.90 0.64

P 0.17 0.13 0.18 0.14 0.25 0.05
Fe 0.06 0.18 0.15 0.12 0.29 0.27

The mineral values, shown in,Table 3, ilndicatesome variations amonoG

the samiiplesof Spartina. Of particular interestis the loss of caleiumiiand
phosphorrus fromthe weatheredstems,and the apparent retentionof iron
in these stemns.
The (lata in Table 4 indicate that growthifaetors of the B vitamin
complex are present in marsh grass, probably in adequate amounts to
TABLE 4. Vitamin content of marshtgrasses expresse(d(as myig.
per 19o0gi. dry weight.
Source of samples is given in Table 2.

Samples
Vitamins 2 3 6
Mature Leaves Mature Leaves YounigLeaves
Thiamine 0.026 0.056 0.037
Riboflavin 0.640 0.725 1.360
Niacin 1.750 2.140 3.520
Pantothenic acid (Total) 1.850 0.768 1.870
B3iotin 0.012 0.010 0.018
Folic acid (Total) 0.288 0.459 0.483
Vitamin B,2 Activity 0.001 0.001 0.002
Pyridoxine 0.418 1.210 0.915
Choline 47.600 61.000 76.500
Inositol 25.600 25.600 43.900

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195] BURKHOLL)ER: NUTTRITIVE VALUE OF SPARTINA GRASS 331

supplv requirementsof certain exacting microorganismsthat may live in

the marsh enivironment. From the amount of growthfactor required for
half-maximumgrowthby the microorganismsemployedin making assays
of these B vitaminis(Barton-Wright1952), it is possible to calculate the
correspondingdilutions of the dried grass in aqueous media, that would
yield equivalent concentrationsfor half-maximumgrowth. These calcu-
lated dilution factors are given in Table 6. Each value represeiitsthe
factorwhich,when multipliedby 100 gm. of grass, would give the grams
of mediumthat would contain sufficient vitamin to permithalf-maximum
growth of heterotrophicmicroorganisms.It is obvious that vitamin-re-
quiringbacteria should findmorethan adequate levels of these compounds,
while growingon or near decomposinggrass in the marsh regions.
TABLE 5. Amninoacid content of marsh yrass expres,sedin per cent (gil. amnlinio
aci(d
per 100 gm. of dry material), and as per cent on the basis of X4 =16. The source of
samples is given in Table 2.

Samiiples
Amiinlo
Acidls 6
Mature Leaves Mature Leaves Young Leaves
Dry Basis N =16 Dry Basis N 16 Dry Basis N = 16

Argiiiiiie 0.214 1.97 0.182 1.61 0.230 1.42

Leucine 0.362 3.33 0.214 1.89 0.307 1.89
Isoleuciiie 0.224 2.07 0.143 1.26 0.252 1.57
Lysine 0.436 4.03 0.769 6.77 0.866 5.35
Methionine 0.026 0.240 0.038 0.337 0.094 0.579
Phenylaline 0.149 1.37 0.128 1.135 0.187 1.15
Tryptophain 0.058 0.534 0.086 0.76 0.124 0.765
Histidine 0.010 0.092 0.028 0.248 0.039 0.241
Valine 0.088 0.710 0.149 1.31 0.197 1.21
Threonine 0.138 1.27 0.256 2.26 0.829 2.03

4 The per cent of nitrogeii oii a dry basis was as follows: No. 2, 1.74; No. 3, 1.81;
No. 6, 2.59.

The presenceof trace amoun-tsof vitaminB12 activityin the grass, as

determinedin the Lactobacillts leichmanniiassay, has been observedalso
in separate assays performedwith the E. coli mutant technique (Burk-
holder and Burkholder1956). It appears possible that cobalamine formed
by microorganisms growingin the water and mud of the region may have
contributedto the assays, either as adhering matter or by absorption of
the vitamininto the tissues of the plants.
Further inspectionof the data in Table 4 suggeststhat adequate levels
of B vitaminsare presentin dried grass to satisfygrowthof the guinea pig,
with the exceptionof thiamineand folic acid (Jukes and Stoksdad 1951,
Mannering 1949). It is, of course, difficultto assess the vitamin require-
mentsof marine animals with our present state of knowledge.
The data in Table 5 indicate that the proportionof amino acids in the

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332 BULLETIN OF THE TORREY BOTANICAL CLUB [VOL. 83

proteinsof Spartina alterniflora,Loisel. is unusual in several respects.If

one compares the amino acid compositionof the leaves of representative
species in the family of grasses (Lugg 1949) with Spartinca,it becomes
apparent that the latter has a relatively low per cent content of such
compoundsas arginine,leucine, isoleucine,methionine,and histidine.The
most strikingdeviation is found in the case of arginine. Accordingto the
data compiledby Lugg (1949), leaves of the Gramineaecontain about 13.7
per cent of arginine in their protein,whereas, in Table 5 only 1.61 per
cent of arginineis reportedfor the proteinof Spartina. Further considera-
tion of the protein compositionof marine fishand shellfish(Walford and
Wilber 1955) indicates that for these marine animals, Spartina proteins
would have somewhatlimited biological value because of their relatively
low contentof arginine,the leuciiles,methionine,phenyrlalan-ine, histidine,

TABLE 6. Calculated dilution factors5 of dried grass (Sample No. 3) IU71iclt when
placed in aqueous media would produce extracts containing an eqnivalent aim?outnt
of each
factor for half maximum growth of typical assay microorganismis.

Vitamins Dilutioin Factor

Thiamine 200
Riboflavini 1,200
Niacin 1,400
Pantothenic acid 3,800
Biotin 10,000
Folic acid 10
Vitamin B12 170
Pyridoxine 240
Choline 800
Inositol 340

5 Dilution Factor (D. F.) =mg. of vitamin in 100 gm. of dried gr8iss/lng.of vitami
in 100 gm. of medium for half-maximumgrowth.

and valine. All of these amino acids of Spartina, when calculated accord-
ing to methodsof Mitchellaild Block (1946), show more than 60 per cent
deficitswhen compared with the correspondingcontentsof amino acids
of beef muscleand fishmLusele(Table 7). Of course,such calectlationsmay
have nutritionalsignificanceonly when "the compositionrepresentsthe
pattern of amino acids which are absorbed into the animal" following
digestiveand otherpreparatoryprocesses (Allison 1949).
If marine animals resemblethe rat and man with regard to require-
mentsfor essential amino acids in normal growthand mainteinaiiee, then
the question arises concerningthe source of adequate proteinfeedstuffs in
the sea. In addition to marsh grass, the second source of primaryfood in
the coastal waters of Georgia is the phytoplanktondiatomsand flagellates.
Accordingto the compileddata of Walfordand Wilber (1955), diatomsare
lower in argininethan marsh grass, but somewhathigherill histidine,and

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19561 BURKHOLDER: NUTRITIVE VALUE OF SPARTINA GRASS 333

about tei timesas high in tryptophan.Althoughthe available data are too

few, still the indicationsare that high quality proteinmay not be formed
by either marsh grass or phytoplankton.There remains the unexplored
possibilitythat microbial conversionmay act like a huge transformerto
step up the potentialvalue of the pool of proteinin the sea. Examination
of the data available fornon-marineLactobacillus,Bacillus and Escherichia
(Lugg 1949) indicates that their per cent compositionof the essential
amino acids is higherthan in Spartina. It appears worthwhileto pursue
furtherstudies on the physiologicalactivities of bacteria that are known
TABLE 7. Deviation(; in per cent of amino acids of No. 3 Spartina sample below or
above the amino acids in proteins of beef, fish and average Gramineae. (Data from
Block, 1945; Lntgg,1949).

Aminiiio
Acids Deviation from Deviation from Deviation fromi
Beef Muscle Fish Muscle average Graniineae

Arginine - 79 - 78 -90
Leucine - 75 - 73 56
Isoleucine - 80 - 79
Lysine -16 -24 + 7
Methioninie - 89 - 90 - 76
Phenylalanine - 77 - 75 - 55
Tryptophane - 41 -41 - 60
Histidine - 91 - 63 - 93
Valine - 77 - 77 - 60
Threonine - 51 - 50 - 24
x-
Per cenlt deviatio= . 100, wvherex is the amino acid of oine protein
-v
(Spartina) and y is the corresponding amino acid of another protein (beef, etc.), the
values of x and y being given in gm. per 100 gm. of protein, based upon N= 16 per ceilt.

to grow upon marsh grass and planktonicresidues in the marine environ-

ment.
SUMMARY

1. Samples of young,and mature,leafy shoots,and weatheredstemsof

the imiarsh grass Spartina alternifloraLoisel. were collected,from marsh
regions along the coast of Georgia, for proximate analyses, and for the
determinationsof ten B vitaminsand ten essential amino acids.
2. Proximate analysis of young Spartina yields values similar to those
whichhave been publishedfor Coastal Bermuda grass. The followingdata
(per cent dry basis) representthe contentsof growingshootsof Spartina:
fat, 2.98; protein,13.24; crude fibre,29.75; ash, 12.83; and nitrogen-free
extract,40.20. The thermsper pound are 1.68.
3. Weatheredshoots,collectedin August fromgrowthof the preceding
year, show loss of theirleaves, and greatlyloweredconteiitof fat, protein,
calcium and phosphorus.
4. The B vitamins,in general,appear to be preseiitiniadle(quiate
amounts

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334 LULLETIN OF THE TORREY BOTANICAL CLUB [\ ()1.. 83.

forgrowthof guinea pigs, as is indicatedby comparisonof the vitaminlevel

in the dried grass and in experimentalrations.The B vitaminsare present
in sufficientamounts for growthof heterotrophicmicroorganisms.
5. The ten essential amino acids, required for growthof the rat aiid
man, constituteabout 15 per cent of the protein of Spartinia. Biological
value of the protein is calculated to be deficientas a nitrogensupply for
marine animals, and it is suggestedthat microbialconversioniof the orass
may act as a transformerto step up the potential value of the pool of
protein in the sea.
BROOKLYN BOTANIC GARDEN
BROOKLYN 25, NEW YORK

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